A Colour Atlas of AVIAN ANATOMY J. McLelland DATOS AGROP. "Bd ; 2 ayeA Colour Atlas of Avian Anatomy John McLelland BVMS, MVSc, PhD, MRCVS Reader in Veterinary Anatomy Department of Preclinical Veterinary Sciences University of Edinburgh Scotland Foreword by: Dr Julian J. Baumel Wolfe Publishing LtdCopyright © John McLelland, 1990 Published by Wolfe Publishing Ltd, 1990 Printed by BPCC Hazell Books Ltd, Aylesbury, England ISBN 07234 15757 All rights reserved. No reproduction, copy or transmission of this, publication may be mace without written permission, No part of this publication may be reproduced, copied or transmitted save ‘with written permission or in accordance with the provisions of the Copyright Act 1956 (as amended). or under the terms of any licence permitting limited copying issued by the Copyright Licensing Agency, 33-34 Alfred Place, London, WCLE 7DP. Any person who does any unauthorised act in relation to this publication may be liable to criminal prosecution and civil claims for damages. ACIP catalogue record for this book is available from the British Library ‘This book is one of the titles in the series of Wolfe Atlases. a series that brings together the world’s largest systematic published collection of diagnostic colour photographs. For a full list of Atlases in the series, plus forthcoming titles and details of ‘our surgical, dental and veterinary Atlases, please write to Wolfe Publishing Ltd, 2-16 Torrington Place, London WC1E 7LT, EnglandContents Foreword Preface Acknowledgements External features and integument Skeleton Digestive system Urogenital system Peritoneum and peritoneal cavities ‘Thorax and neck Respiratory system Central nervous system Radiographic anatomy Further reading Index 95, 120 12 123 124Foreword The focus and expertise of many of us whose research deals with avian anatomy is rather narrow. Relatively few individuals are widely knowledgeable generalists in avian anatomy. The author of this book is one of the latter who not only has done much original research on the structure of birds, but has in his background the experience gained as the author of a number of critically acclaimed books on avian anatomy. Furthermore, as one of the principals of the International Committee on Avian Anatomical Nomenclature (ICAAN), Dr McLelland is responsible for the development of a standardized terminology for the anatomical parts of birds. ‘Most anatomical illustrations are drawings, usually highly schematic. Photographic atlases have the advantage of portraying not idealized interpretations, but images of the actual structures themsclves. Some photographic anatomical atlases are disappointing, their illustrations lacking definition and perspective, certainly not the case in this book. The author himself has produced an outstanding series of colour photographs that overcomes the intrinsic difficulties of photographing anatomical material, He has presented a balanced array of illustrations that not only introduce the structure of birds to novices, but are also so distinctive and fresh that they are instructive and enlightening to those of us who deal with avian morphology routinely. ‘This atlas is indeed a worthy accomplishment and an important contribution to avian science. Dr Julian J. Baumel, Professor of Anatomy, School of Medicine, Creighton University, Omaha, Nebraska General Chairman of [CAANPreface There is considerable interest today in birds whether as food producers, as models for biological research, or simply as objects of pleasure in aviaries and in the wild. This colour atlas provides an attractive and easily assimilated introduction to how birds are constructed. The book is aimed primarily at those engaged in university studies, veterinary practice, meat inspection and research, but itis also hoped that it will be of interest to ornithologists and bird-watchers. The book is concerned principally with the economically important species but where their anatomy significantly differs, the more common cage birds are dealt with. The choice of material is determined by the author's experience over the years in research and in university teaching at both undergraduate and postgraduate levels, and reflects the needs of veterinary surgeons as well as covering topics of general biological interest. Most of the 309 photographs are of fresh, unfixed material: 49 have been taken with the aid of a dissecting microscope. Light micrographs and scanning electron micrographs have been included when they enhance understanding. Scale bars are used only when they are thought to be of positive value. The photographs are accompanied by a short text which aims both to identify the various features and also to explain their function. For additional information the reader is directed to the ‘Further reading’ list at the end of the book. The book is planned basically around certain body systems including the integument and skeleton, and the digestive, urogenital and respiratory tracts Lymphoid tissue, blood vessels and nerves are dealt with as they occur in association with these systems. In some instances a straight topographical approach is taken, as in the description of the peritoneum and peritoneal cavities ‘The names in the text of the common birds are as follows: duck, domestic forms of Anas platyrhyrchos; goose, domestic forms of Anser anser: chicken, domestic forms of Guilus gallus; turkey, domestic forms of Meleagris gallopavo: quail. domestic forms of Cotwmnix; pigeon, domestic forms of Columba livia, Budgerigar, Melopsitiacus undulatus; Canary, Serinus canaria.Acknowledgements 1am very grateful to the following for allowing me to use their photographs: Dr M.A. Abdalla of the Department of Anatomy. Faculty of Veterinary Science, University of Khartoum, Sudan for 287; Dr E.M. Abdel-Magied of the Department of Anatomy, Faculty of Veterinary Science. University of Khartoum, Sudan for 232; Dr G-H.A. Borst for 302; Mr JE. Cooper and the Royal College of Surgeons of England and Dr J. Samour and the Zoological Society of London for 184; Professor A.S. King and the Department of Veterinary Preclinical Sciences. University of Liverpool for 129, 130, 164, 170, 171. 183. 186. 201, 288; Dr P.E. Lake for 202, 203; Dr A.M. Lucas for 288. 289, 290. 304; Dr J.N. Maina of the Department of Veterinary Anatomy, University of Nairobi, Kenya for 285; Dr M.D. Purton of the Department of Veterinary Anatomy, University of Glasgow for 11, 18, 229, 266, 267, 297, 300; Mr C.J. Randall of the Ministry of Agriculture, Fisheries and Food, Lasswade, Penicuik for 114. 162, 166; Dr S.E. Solomon of the Department of Veterinary Anatomy, University of Glasgow for 167, 172, 173, 176, 177, 178; Dr M.K. Vidyadaran of the Faculty of Veterinary Medicine and Animal Science, Universiti Pertanian Malaysia, Serdang, Selangor, Malaysia for 282, I would like to thank the following publishers for allowing me to reproduce work in which they hold the copyright: Academic Press, London; Birkhauser Verlag, Basel; Cambridge University Press; Elsevier Science Publishers B.V., Amsterdam. I am greatly indebted to Mr and Mrs E. Denby for enabling me to take the photograph for 25 and to Fenton Bams (Scotland) Ltd, North Berwick for permission to take the Photographs for 4, 22, 23 and 24 Almost two-thirds of the photography was performed by Fiona Manson and I am greatly indebted to her for the endless time and trouble she spent on the project. Other photographs were taken by Colin MacFarlane, Adnan Mahdi, Stephen Mitchell, Derek Penman, Kenneth Thomson and Colin Warwick to whom I express my sincere thanks. The photographs with the dissecting microscope and the remaining photomicrographs were taken by me with the skilled assistance of Gordon Goodall. 1 am also indebted to John Strathearn and Ian Warwick for their help in obtaining specimens. Finally, I would like to thank the staff of Wolfe Publishing Limited for their assistance in producing this book.External features and integument 1.4 Regions of the body. (a) Cranium; (b) face; (c) Oral region: (2) nasal region; (3) orbital region; (4) neck: (d) back: (e) rump: (f) tail; (g) side; (h) brea- ear region; (5) forehead; (6) crown; (7) occiput; (8) st; (i) belly; (j) undertail or erissum: (k) arm; (I) chin; (9) throat; (10) cheek; (11) nape of neck; (12) forearm; (m) hand: (n) thigh; (0) leg; (p) foot. (1) jugulum. 2 Wing of an adult pigeon, 1 Adult male duck. 3 Leg of an adult chicken, 4 Head of an adult female turkey.5 External features of an adult male duck, The over- all shape of the body is modified by the externally visible plumage. the contour feathers, The main ty= es of contour feather are the flight feathers of the ‘wing (a) and tail (b), and the general contour feat. ers (c) of the head, trunk and limbs. The contour feathers overlie the other feather types including the scmiplumes, down feathers, bristles and filo- plumes. In some regions such as the bill (d), scales (e) and claws (f) the skin is much more heavily keratinized than elsewhere and is without feathers, 7-10 Bill of @ duck. Whilst the keratin covering most of the bill in the duck is soft and leathery. at the rostral end there isa hardy horny plate. the nail (a), supported by a thickened region of bone (b). With the mouth closed the upper nail lies almost at right angles 10 the lower nail. Running through ‘each nail are numerous canals which open on the ‘occlusal surface of the nail. The canals contain der mal papillae carrying mechanoreceptor nerve end- ings. On the bone below each nail are pits (arrow) which also contain dermis with sensory nerve end: 0 6 Beak of an adult chicken. The beak or bill inclu- des parts of the skeleton of the upper and lower jaws and the hard horny sheaths or rhamphothecae covering them, The median dorsal border of the upper or maxillary rhamphotheca is the culmen (2). The median ventral border of the lower or mandibular rhamphotheca is the gonys (b). The ccutting edges of the rhamphothecae are the tomia, (c). The maxillary rhamphotheca is perforated by the paired nostrils ings. The upper and lower nails and their associated dermis form complex sensory organs. the upper and lower bill tip organs. which are important in the sensory discrimination of food. A well-developed Dill tip organ is also present in parrots. Along the ‘edges of the rhamphothecae in the duck are blade= like lamellae (c), one series on the inner margin of the upper bill and two series on the outer margin of the lower bill, which with the bristles (d) of the tongue are used for straining food.7 Upper horny bill of an adult duck. 9 Bony part of the upper bill of an adult duck. 10 Lamelia of the lower bill of an adult duck11-14 Beaks. In the budgerigar the upper beak is stout, sharp-tipped and strongly curved and closes ‘over and hides most of the small, blunt lower bes TThe tips of the upper and lower beaks are composed ‘of much harder horn than elsewhere. The sharp, pointed upper beak of psittacines may have to be blunted to prevent damage to the owner. At the base of the upper beak in the budgerigar is the swollen, sensitive cere (arrow). In young birds of both sexes the cere is a greyish-blue colour but after 3 months it usually turns blue in the cock (11) and light brown in the hen (12), However, in some coloured varieties of budgerigar, such as, lutino mutations, the appearance of the cere in the male and female is very similar and therefore sexing is not easy. Budgerigars up to the age of 3 months are sometimes called "barheads’, because the plumage 11 Beak of a male budgerigar. cof the erown and forehead! as far as the cere is eros- sed by distinct dark bands similar to those on the back of the head of the birds in the photographs ‘These marks on the crown and forehead are lost {ter the first moult. In the pigeon (13) there is also. cere (arrow) at the base of the beak but unlike that ofthe budgerigar itis white and lies behind the nostrils. The beak of the canary (14) has a conical shape typical of seed-cating passerine species. Avi- calturists classify cage birds into hardbills which include canaries, budgerigars and most parrots and softbills which include mynahs, starlings and thrushes. The beak plays an important part in pre~ hersion and in the physical preparation of food. and in some species Such as parrots itis even used aan aid to locomotion. 12 Beak ofa female budgerigar. 13 Beak of a pigeon. 14 Beak of a canary.1S Egg tooth of a day-old chicken, The ege tooth (arrow) is a conical protuberance of hard keratin which develops before hatching and assists the chick in pipping or breaking out of the shell. Afte hatching itis lost within 1-2 days. The shape of the beak in chicks of different species is remarkably similar, despite the fact that the adult beak shows such a wide interspeci ion in its appea 16-18 Feet. The foot of most avian species like that of the canary and duck has four toes, three toes pointing forwards and one toe (digit I) pointi backwards. This arrangement in the canary is an adaptation for perching. In ducks the three forward pointing toes are webbed for swimming. The foot of the budgerigar and parrots has two toes (digits IL and Ill) pointing forwards and two toes (digits | and IV) pointing backwards and is an adaptation for climbing and grasping. The terminal phalanges are cnclosed by the claws which, in caged birds, are sometimes not subjected to the normal wear and may therefore have to be clipped. Clipping the claw is also carried out to obtain a blood sample. 16 Foot of acanary. 18 Foot of a budgerigar. 17 Foot of a duck.19 Foot of an adult male chicken. Dorsal view. The chicken has a typical avian foot with three toes pointing forwards and one toe pointing backwards. On the caudomedial surface of the tarsometatarsal region of the male chicken, pheasant, grouse and turkey is a well-developed heavily keratinized spur (arrow) with a bony base (see 101). The spur grows, by about I em every year until a maximum length of approximately 6 cm is reached. A cock’s spur should be kept trimmed to a length of about 13 mm to avoid damage to the hens during mating. The spurs usually very small or absent in females (20). 4 : if . 20 Foot of an adult female chicken. Plantar view. ‘The joints of the foot are supported on their plan- tar sirfaces by elevated pacls consisting of thickened skin covering a core of subcutaneous adipose tissue In the chicken there isa single metatarsal pad (2) at the metatarsophalangeal joints and several digital pads (b) on each of the main digits. The spur (arrow) In the female is very reduced. In the slaughter plant the feet are cut off anywhere in the tarsometatarsal region, although in some countries they are removed at the hock joint21 Ornamental appendages on the head of an adult male chicken. Ornamental appendages are especi- ally well-developed in male birds and include the mb (a), wattles (b) and rieti (c). They are disten- le structures characterized by a very thickened and vascular dermis containing, numerous arterio- venous anastomoses. The colour of the comb may be used to ascertain the well-being of a bird. A bright red, full comb is a clear indication of onset of layin a pullet. In the poultry slaughter plant the comb reflex is used to determine if a bird is conci- us or not. Dubbing or removal of the comb in day ‘old chicks is sometimes carried out to avoid trauma tization or frostbite. The wattles may also be cropped in chicks to avoid frostbite. The edges of the wattles are sometimes used as a site for intradermal injee- tions. The lobes (d) hang down below the external ear openings and are cither red or white depending upon their vascularity22-24 Ornamental appendages on the head of the adult male turkey. The snood or frontal process (a) of the adult male turkey is extensible like the comb of the chicken, its size varying with the emotional state of the bird. Unlike the comb, however, the dermis of the snood has much smooth muscle but c tissuc. Removal of the snood or desnoo- carried out in chicks to prevent trauma tion, A dewlap (b) arises from the region of the throat. The head (c) and upper neck (d) skin of the turkey is carunculate, In intensively raised turkeys and chickens, part of the upper beak is usually cut off as in the photographs to prevent feather pecking and cannibalism, Because the beak is richly inner- vated. it should not be trimmed more than one-third of the distance from the tip to the nostrils, 23 Head of an adult mate turkey. 24 Head of an adult male turkey.25 Prevention of cannibalism in a pheasant. Methods other than debeaking are sometimes used to prev cent cannabilism. In pheasants, for example, a plas tic “bit” which extends below the palate from one nostril to the other may be used. The size of the it’ is adjusted as the bird grows. 26 Skin of the breast region of an adult chicken, Except in obviously horny sites the skin of birds i, thin, dry and white to yellowish-pink in colour. It also appears to be relatively avascular compared to the skin of mammals, because it docs not bleed as profuscly if cut. However, because the skin i only loosely attached to the underlying: musculature. any bleeding which does occur can extend under the skin for considerable distances and result in un- sightly bruises. Inthe poultry slaughter plant these blemished areas have to be trimmed away from the carcass, The subcutaneous layer contains fat which may be abundant. and striated muscle which acts to regulate the tension of the skin. Because the skin is relatively inelastic. subcutaneous injections are often accompanied by leakage ofthe injection uid through the puncture point of the needle. The best sites for subcutaneous injections appear to be the dorsal side of the dase of the neck, the breast region and the groin. Tn most species the skin in the breast region is modified during brooding to form an incubation (brood) paich or patches. Here the dermis becomes thickened and very vascular and the feathers are lost. At these areas the body heat of the bird is trans- ferred to the incubating eggs. Incubation patches ‘may occur only in one sex (generally the female) but in some species, including the pigeon. they develop in both sexes, They are absent in ducks ‘and geese: they keep their eggs warm in the nest by covering them with plucked down feathe27 Uropygial gland of an adult duek. The bilobed uropyaial gland or preen gland is the main skin aland of birds (there are no sweat glands and only a few sparsely distributed sebaceous glands) and lies dorsal to the caudal vertebrae near the tip of the tail (see also 267). Its holocrine sebaceous secretion, sebum. is smeared over the feathers during preen- ing, and by forming a filn of fat helps to keep the thers waterproof and to prevent their keratin drying out and becoming brittle. Sebum also inhibits the growth of micro-organisms so that skin infect- ions in birds are relatively rare. In addition to the preen gland, sebum is also secreted by the epider- mal cells of the skin. The uropyzial gland is well developed in the duck and budgerigar but is absent in many parrots and in some strains of domestic pigeons, 28 Uropygial gland papilla in an adult chicken. The ducts of the urepygfal gland open on a median nipple like papilla (arrow) which is situated dorsally near the tip of the tail29 and 30 Contour feathers. ‘The contour feathers ive the body its form and can be divided into gene- ‘al feathers of the head, trunk and limbs, and flight feathers. The shaft of a contour feather is composed of the calamus or quill (a) which is implanted in the feather follicle, and the rachis (b). On either side of the rachis is a row of fine branches, the barbs (c) which together form a vane (d). Arising from the barbs are branches, the barbules. The darbules in the distal part of the general contour feather and ‘over most of the flight feather are hooked together 29 General contour feather of an adult chicken. so that each vane is flat and firm. In the proximal parts of the feathers the barbules are not hooked together so that the vanes are fluffy. Contour fea~ thers form en almost continuous insulating layer and help to maintain the high body temperature of 105-107°F (40-41°C). They are also important in flight. in display during fighting and courtship, and a8 nest material. In waterbirds contour feathers trap air and enabie the bid to float by maintaining buoyancy 30 Flight feather of adult chicken,31 and 32 Calamus and feather follicle. At the tip Of the calamus is an opening. the proximal umbili- cus (a). The proximal part of the calamus lies in a tubular invagination of the skin called the follicle (b). Projecting for a short distance through the proximal umbilicus is the dermis at the base of the follicle, which becomes continuous in the cavity of the calamus with a small mound of pulp called the to the nest feather. The calamus is hollow and tans. lucent and is divided into chambers by epidermal partitions, the pulp caps (arrow). Attaching at each end by elastic tendons to the sides of the follicles of the contour and other feather types, are smooth muscle bundles which are responsible for feather fluffing to conserve heat. Because the muscles often form extensive parallelogram-like networks. ermal papilla. Covering this very vascular papilla are live epidermal cells which after moulting give rise. along with the cells at the base of the follicle. wide groups of feathers can be acted on at the same time. Calamus of a contour feather of an adult chicken. 32 Embedded part of a contour feather of an adult chicken. 33 Undersurface of the flight feather of am adult ‘chicken. At the junction of the rachis (a) and cala- mus (b) a groove (arrow) on the surface of the shaft terminates in an opening (c) called the distal umbili- cus. The distal umbilicus communicates with the y of the calamus. There is often a small after er (83) attached to the rim of the distal umbili 20BM and 38 Vane of a flight feather. The rachis (a). ves off on each side at about 45° to the main shaft a series of stiff branches, the barbs (b). Arising from the barbs at about 43° are finer branches, the barbules (c,d). The barbules of adjacent barbs cross each other at 90°. The distal barbules (c) are directed towards the free tip of the feather and carry hooks. They interlock with the proximal bar- bules (d) which are directed towards the attached end of the feather. In this manner a firm and flat vane is formed. If the barbules are unhooked when the vane is split, the bird re-engages them during preening by pulling the vane through its beak in a direction parallel to the barbs. 3M Scanning electron micrograph of the vane of a chicken Might feather. (Scale, 100 jum.) 36 and 37 Pterylosis. Contour feathers are not evenly distributed over the body but are arranged n tracts called pterylae which are separated by relatively bare spaces or apteria. The arrangement of the pterylae is characteristic for each species. Within the apteria are usually down feathers and semiplumes. Figure 36 shows two pterylae (a) in 38 Scanning electron micrograph of the vane of a chicken fight feather. (Seale, 5 jm.) the breast region, the feathers of which in 37 have been pushed aside to reveal an apterium (b). Within the dermis of apteria are smooth muscle bundies which tense the skin. Whenever possible surgical incisions should be made in apteria rather than in prerylae. 36 Pterylosis of an adult chicken. 37 Pterylosis of an adult chicken.39 and 40 Wing feathers of an adult duck. The feathers of the wing consist of flight feathers or remiges and coverts or tectrices. The remiges are cither primary feathers (a) attaching to the meta- carpal and phalangeal bones or seconcary feathers () attaching to the ulna. In the duck there are 10 well-developed primary remiges which are numb- ered in a medial to lateral direction and 18 second- ‘ary remiges which are numbered in a lateral to medial direction. All the remiges overlap in the same way, the trailing edge of one feather being overlain by the leading edge of the next feather. Because the bsses of the primary and secondary feathers are interconnected by the elastic inter- remigial ligament, the unfolding of the remiges is coordinated. All the secondary feathers and the inner primaries are relatively movable. The outer 39 Wing feathers of an adult duck, Dorsal view 38 Pterylosis of plucked 8-week-old chicken. Ven- tral view. In the chicken there are about 70 named pterylae or tracts: (a) ventral cervical tract; (b) pec toral tract; (c) sternal tract; (d) abdominal tract: (e) crural tract, (I) Ventral cervical apterium: (2) pectoral apterium: (3) sternal apterium: (4) crural apterium primaries, in contrast, ere more firmly fixed. In many species a gap of diastema is present in the secondaries. The flight feathers are covered proxi mally by rows of wing covers (c). To prevent escape in parrots and other large cage birds or to reduce the mobility of an aggressive cock during the breed ing period. trimming of the flight feathers in one wing is recommended. Usually all the primary and secondary feathers are cut except the outermost two and the innermost three. For cosmetic reasons the cut is made just below the level of the coverts so that the trimmed ends of the feathers are not seen, With one wing trimmed in this way the bird unbalanced and cannot maintain height. Because the feathers will be replaced at the next moult, trimming has to be done annually 40 Wing feathers of an adult duck. Ventral view.41 Leading edge of a chicken wing. Ventral view. The triangular fold of skin that forms the leading edge of the wing between the shoulder and carpal joints isthe propatagium (a). In its cranial border is ial (b). The propata- _glum serves to increase the surface area of the wing ‘and is acted on by a number of muscles. Removal of @ triangular portion of the propatagium has sometimes been carried out to make cage birds flightless. In the ringing of ducks wing clips may be attached to the propatagium. The ulnar vein (c) is frequently used for intravenous injections. Other veins which can be used for injections are shown in 100, 229 and 230. 42 Extensor carpi radialis tendon of an adult chick- en. The short tendon (arrow) of the extensor carpi radialis muscle stretches across the cranial surface of the carpal joint and is sometimes cut in one wing to prevent a bird from flying. 43 Alula or bastard wing of a budgerigar: (2) feath- ers of alula; (b) primary flight feathers: (c) secondary flight feathers; (d) coverts. The alular digit in the budgerigar curries four feathers (between two and seven in birds generally) which form the alula. The alula when pulled forward gives rise to a slot which ensures that a smooth stream of air is present on the upper surface of the wing for lift when the bird's, speed falls too low or the angle of attack is raised above 20°,44 Flight feathers of the tail of an adult pigeon. The ht feathers of the tail, the rectrices, are always jaterally paired. In the pigeon there are six feath- ers on each side, The recirices are covered dorsally and ventrally by small tail coverts, Because of the free nature of the caudal vertebrae (see 79), the tail, is very movable and assists the wing in steering and braking, as well as being important when depressed and spread at low speeds as an additional wing area, 45 Semiplume of an adult chicken. The vane is fluffy and the rachis is longer than the longest barb. Semiplumes occur along the margins of the contour feather tracts and singly within the tracts, They are hidden by the contour feathers and assist im insula ting the Body 46 Structure of a semiplume of an adult chicken. Originating from the long rachis (a) are barbs (b). each barb gi fbules (c). The barbules are not hooked together so that the vane is fluffy (Scale, 5 mm),47 Adult down feather of a chicken. The vane is fluffy and the rachis is cither absent or shorter than the longest barb. Adult down feathers may be res- tricted to apteria, ay in chickens, evenly distributed over the body as in ducks and parrots, or sparsely distributed in apteria or entirely abse and passerines. Down feathers ulatory function and in waterbirds they assist con. tour feathers in trapping air and maintaining buoy 48 Natal down of a day-old chick. In newly-natched ickens and ducks the down forms a dense layer ‘over the whole body, but in pigeons and passerines it is restricted to the contour feather tracts. Natal down feathers differ from adult down feathers in generally lacking barbules on the tips of the central ancy. Another type of down feather which occurs in some species, including parrots and pigeons (but not chickens), is the powder down feather which liberates a white waxy powder of keratin granules, ‘The granules probably play a part in water-proofing ers. Because of their abundance in ducks, down and other fine feathers are removed in the poultry slaughter plant by a waxing process.50 and 51 Filoplumes. Following plucking, heir like feathers called filoplumes usually remain on the carcass. They are characterized by & slender 49 Bristle of an adult chicken. Bristle feathers typi cally consist of a stiff rachis and a few barbs at their proximal end. The feather particular type of bristle called a semibristle in which the barbs arise from most of the length of the rachis, Bristles occur almost exclusively in the head iand neck and include the so-called ‘eye-lashes’ (60) and the narial bristles around the nostrils. Because the follicles of the bristles are surrounded by nume~ rous sensory Herbst corpuscles, it has been sugges ted that bristles are tactile sensory structures ana logous to the whiskers of cats. (Seale, J mm) stalk and a short tuft of barbs or barbules at their tip, ‘50 Filoplume of an adult chicken. 26 S1 Filoplume of an adult chicken. (Seale, 1 mm) 52 Filoplume of an adult chicken. Filoplumes (arrow) never occur alone but always accompany other feathers which are generally semiplumes anc contour feathers, Unlike other feather types they are not associated with smooth muscle bundles in the dermis. Within and around the walls of the filoplume follicles are numerous nerve endings: it has been suggested that the filoplumes are responsible for a proprioceptive sensory input ‘which ensures that the feathers they are associated with are correctly positioned. In the poultry slaughter plant the filoplumes are left on the carcass after the birds have passed through the plucking machine; therefore they have 19 be removed by a process of singeing.S3.and $4 Afterfeather. The afterfeather (arrow) is a small feather attaching to the underside of a con tour feather or semiplume at the rim of the distal umbilicus. Afterfeathers range from a downy tuft toa highly organized feather with well-developed 53 General contour feather and afterfeather of an adult chicken, vanes similar to the distal part of contour feathers. In the afterfeather illustrated here the barbules are not hooked together so that the vane is entirely fluffy. It is likely that afterfeathers have a thermal insulatory role, 54 ANterfeather of §3 enlarged.‘55 Wing feathers of an adult budgerigar. Feather colour and moulting. In the budgerigar there are 10 primary remiges (a) and 11 secondary remiges (b) Feather colours are either pigmentary or structural, ‘or a combination of both. Colours which are pigmentary in origin are due to pigments in the feathers, the principle ones being melanin, carotenoids and porphyrins. Colours which are structural in origin are due to the form of the feather surface and may originate from interference which produces iridescence 2s in starlings, or the scattering of light which results in noniridescent colours such as the blue of the budgerigar, resulting fi Green is an example of a colour mm both pigment and structure and is a combination of yellow carotenoids and. Tyndall blue scattering. The colour pattern of feathers frequently shows sexual dimorphism and in many species the male is more highly coloured than the female. This does not apply, however, to the jority of small cage birds which are not sexually dimorphic. Variations in colour patterning may also be related to the season of the year or the age of the bird Moulting of feathers usually occurs once a year and generally after the breeding season in summer or autumn, Passerines typically moult sometime between May and December, the moult lasting between 5 and 8 weeks. Psittacine species can ‘moult throughout the year. some birds appearing to moult continuously. The annual moult of adult pigeons is usually in August or September. In the chicken there is normally a complete moult in the auturan. The replacement of flight feathers in most species is carried out in a certain sequence, so that the bird retains its ability to fly. In ducks, however, all the large fight feathers are lost simultaneously. ‘so that flight for a period is impossible.‘86-59 Ear coverts and the ear. There is no auricle or ear flap in birés so that the external ear opening is flush with the surface of the head, It is covered by a cirle of contour feathers, the ear coverts, The coverts (a) on the rostroventral rim of the opening are long and directed dorsocaudally. Those (b) on the remainder of the rim are short and vertically orientated, so that they form a funnel. The rostral feathers impinge on the tips of the vertical feathers and are arranged like a screen keeping out foreign objects. The passage of sound, however, is not obstructed because the barbs of the ear coverts ere further apart than in other contour feathers. In the chicken the opening of the external ear (c) is Greular and about 5 mm in diameter. From the opening the external ear canal (d) extends downwards and inwards to the _ tympanic membrane (¢) which forms the internal boundary of the external ear. The walls of the external ear canal contain holocrine glands. The tympanic membrane stretches across the rostrodorsal part of the tympanic cavity (f). Vibrations of the tympanic ‘membrane are transmitted to the inner ear by the bony columella, the only ear ossicle. The medial footplate of the columella is implanted in the vestibular window (1), The cochlear window (2) is closed by a thin membrane. Opening on the floor of the tympanic cavity is the pharyngo-tympanic tube which connects the middle ear to the pharynx: Via the infundibutar cleft (see 102). 56 Ear coverts of an adult chicken. 57 Ear coverts of an adult chicken displaced rostrally. 58 External ear canal and tympanic membrane of a 10-week-old chicken. Fixed preparation. (Scale. Imm) ‘59 Right tympanic cavity of an adult duck. (Seale, Timm)oo 60 and 61 Eyelids and eyeball. The upper eyelid (a) is relatively immobile and only closes on sleep. The lower lid (b) is much thinner and more movable. With the eyelids open only a very small part of the so that a true indication of the enormous size of the eyeball shown in 61 is not eiven. The large size of the avian eye is related to igh visual capacity. In aliricial species, including budgerigars and passeriforms, the eyelids losed for ashort time after hatching. The small eye is visible eda. (60 Eyelids of an adult chicken, 0 feathers (arrow in 60) at the margins of the lids are bristles, The transparent nictitating membrane or third eyelid (c) lies on the nasal side of the orbit and is moved across the eye in blinking by two striated muscles. Its sweeping movement helps to distribute the secretion of the gland of the membrane, During the electroplectic fit following stunning in the poultry slaughter plant, this corneal reflex disappears. The position of the ring of bony scleral ossicles (d) is seen in 61 (see also 65), 61 Eyeball of an adult chicken. (62 Nasal gland of an adult duck. The nasal gland, (arrow) lies dorsal to the orbit and hasan osmoregulatory function in marine species whigh enables these birds to drink sea water, a 3 per cent saline solution. The gland opens by ducts into the rostral part of the nasal cavity. Within the gland a counter-current system exists between the blood and the secretory cells. The gland is also ‘osmoregulatory in some desert species and in terrestrial carnivores feeding on a very salty diet. It is also present, but non-functional, in many other terrestrial species, including the chicken.63 Gland of the left nictitating membrane of an adult duck. Caudal view. This large gland (arrow) es on the caudomedial aspect of the eyeball. Its secretion flows onto the surface of the eye below the nictitating membrane. Within this gland and the nasal gland are numerous plasma cells which help to protect the eye against infection. (Scale 3 mm) 64 Right lacrimal gland of an adult duck. The lacrimal gland is much smaller than the gland of the nictitating membrane and lies atthe lateral canthus attached to the rim of the orbit. It drains into the conjunctival sac. The secretion of both the lacrimal gland and the gland of the nictitating membrane Passes via two ostia at the medial commissure of the eyelids into the nasolacrimal duct which opens into the nasal cavity below the middle nasal concha (see 237) 65 Bony orbit and scleral ossicles of an adult chicken. The bony orbit is exceptionally large and is incomplete. The cyeball almost totaly fills the orbit, so that eye movements in birds are small up for by the great mobility of the head and very flexible neck. Because of the great size of the eyeballs, the right and left orbits are separated by a very thin interorbital septum (a) The sclera in the region of the cornea is reinforced by a number of small bones. the scleral ossicles (b) which strengthen the eyeball and give attachment 10 the striated cilia les of accommodation ‘see also 61). The remainder of the sclera is roughened by hysline cartilage666 and 67 Pecten of the chicken eye. The pecten is an extremely vascular pleated structure which projects from the retina into the vitreous body. Its base rests on the optic dise while distally the pleats are held together by a bridge which is firmly 66 4 h 66 Pecten of the eye of day-old-chicken. Fixed reparation. (Scale, I mm) attached to the vitreous body. Its black colour is Gue to numerous melanin pigment cells. The function of the pecten appears to be 10 provide nutrition by diffusion through the vitreous body to the avascular retina 67 Pecten of the eye of un adult chicken. Fixed preparation (Scale, T mm)Skeleton 68 Skeleton of an adult chicken. Compared with ‘mammals the skeleton of birds is lighter and has a higher calcium phosphate composition. Its characteristic features include fusion of most of the trunk vertebrae, extreme mobility of the cervical vertebral chain, fusion of the pelvic to the vertebral column and the absenk pelvic symphysis, a prominent steraum, a strong pectoral gitdle, aad simplification of the skeleton ‘of the limbs by the fusion or elimination of bon 69 Skull of an adult duck. Lateral view Characteristic features of the avian skull include the vaulted bulbous brain case (A), the large bony orbits (B), and the beak-shaped pyramidal face (C). The boundaries of individual bones are usually difficult 10 identify because the sutures between many of them’ are lost soon afte hatching: (a) premaxilla; (b) maxilla; () bony external naris; (d) nasal bone; (e) frontal bone: £) lacrimal bone; (g) parietal bone: (h) occipital bone; (i) ethmoid bone (interorbital septum); (j) jugal arch; (k) vomer bone; (I) palatine bone: (m) pterygoid bone; (n) quadrate bone; (0) mandible. To increase the size of the gape, the upper jaw is raised at the junction (arrow) of the cranium with the premaxillary and nasal bones where the bone is thin and flexible and the interorbital septum is absent7” 70 Cranial kinesis in the skull of an adult duck. Lateral view. Cranial kinesis is movement of the upper jaw in relation to the brain case. Because the quadrate bones articulate with the brain case. 71 Skull of @ parakeet, In psittacine species, including the budgerigar, prokinesis is highly developed, the craniofacial elastic zone between the nasal and frontal bones being converted into a synovial hinge joint (arrow). 72 Skull of an adult duck. Ventral view: (a) premaxilla; (b) maxilla; (c) vomer; (d) sphenoid bone; (e) occipital bone; (f) foramen magnum g) palatine bone; (h) plerygoid bone: (i) jugal arch; (j) quadrate bone; (k) mandible. The internal nares (arrows) lie between the palatine bones and the vomer pterygoid bones, jugal arches, and mandible they are pivoting points in cranial kinesis. The particular type of cranial kinesis which occurs in chickens, ducks and known a prokinesis. In the process of raising the upper jaw, depression (1) of the mandible (a) rotates (2) the quadrate bones (b) rostrally and pushes (3) the pterygoid (c) and palatine (4) bones and the jugal arches (e) forward. This results in the upper jaw swinging forward and upwards (4) at the craniofacial elastic zone (f) where the bone is thin and flexible. Coaversely caudal rotation of the quadrates lowers the upper jaw. In many species, including pigeons, the elevation of the upper jaw ‘occurs rostral to the junction of the jaw and brain case and is known as raynchokinesis. In contrast to prokinesis, it involves only the rostral part of73 Skull of an adult duck. Dorsal view: (a! premaxilla; (b) maxilla; (¢) bony external naris (6) nasal bone; (c) prefrontal bone; (f) frontal bone: (g) parietal bone; (h) lacrimal bone; (i jugal arch; ()) pterygoid bone: (k) quadrate bone e craniofacial elastic zone (arrow) lies between the cranium and the nasal and premaxillary bones. 74 Skull of an adult duck. Caudal view: (a) occipital bone: (b) sphenoié bone. The occipital bone encloses the foramen magnum (1). A characteristic feature of the avian skull is the single occipital condyle (2), which makes the atlanto-occipital joint a much more mobile culation than in mammals and allows considerable rotation of the head, Pneumatization of the skull. Some of the skull bones, like the frontal bone of the passeriform species in the photograph, are formed dy two thin plates of compact bone joined together by 2 network of delicate spicules enclosing air spaces continuous with the tympanic and nasal cavities Pneumatization and the concomitant reduction of the weight of the skull is probably an adaptation for flight.7 6 76 18 76 The S-shaped cervical region of the vertebral column of an adult chicken, Left lateral view Unlike in mammals the number of cervical vertebrae varies with the length of the neck. In the chicken there are usually 16 vertebrae, The joints between the vertebrae are synovial and the articular surfaces of the bodies are saddle-shaped. so that this region of the verte extremely flexible, The small r (arrow) articulates with a single occipital condyle. so that the atlanto-occipital joint is exeeptionally mobile. The great flexibility of the vertebral column and the mobility of the atlanto-oceipital joint allows the beak to be used for a wide variety Of tasks, which in mammals would be carried out by the forelimbs, 77 Cervical and thoracic vertebrae of an adult chicken. Left lateral view: (a) cervical vertebrae: (b) notarium; (€) fourth’ thoracic vertebra: (d) scapula; (@) humerus. The notarium is formed by the last cervical vertebra and the first three thoracic vertebrae, the fusion between the vertebrae in the chicken starting at about 4 months of age. The notarium along with the synsacrum gives the spine great rigidity. The vertebrae in the duck and goose remain freely able 78 Synsacrum and pelvis of an adult chicke Dorsal view: (a) iium; (b) ischium; (c) pubis; (d) synsacrum; (e) free caudal vertebrae. The synsacrum consists of fused thoracic, lumbar sacral and caudal vertebrae, the fusion in the chicken starting at about 7 weeks. The synsacrum. is itself fused to the ilium. The antitrochanteric process (arrow) of the pelvic bone is covered by cartilage and articulates with the neck and trochanter of the femur. The joint aets to limit abduction of the limb.79 Caudal part of the vertebral column of an adult. chicken. Right lateral view: (a) synsacrum: (b) ree caudal vertebrac; (c) pygostyle; (d) ilium; (e) schium. In the chicken there are six free caudal vertebrae which permit movement of the tail. The Pygostyle consists of between four and six fused audal vertebrae and provides attachment for the snnermost tail feathers. 80 Trunk skeleton of an adult chicken. Lateral view: (a) cervical vertebrae; (b) notarium; (c) fourth thoracic vertebra; (d) free caudal vertebrae; (e) pygostyle; (f) vertebral rib; (2) sternal rib; (h) sternum; (i) clavicle; (j) coracoid bone; (k) scapula; (I) humerus; (m)’pelvis; (1) femur; (0) tibia and fibula, In the chicken there are five or six pairs of ribs, each rib consisting of a vertebral par! which has @ double articulation with the vertebral column and a stemal part which articulates with the sternum. The uncinate processes (1) of the vertebral ribs provide attachments for muscles and ligaments and strengthen the thoracic wall. The bony sternal ribs correspond to the costal cartilages of mammals and provide support for the thorax during the downstroke of the wing. The vertebral and sternal ribs articulate at_a movable cartilaginous joint. The last few cervical vertebrae articulate with floating ribs (2) which lack a sternal component, The articulations between the thoracic ribs and the vertebral column and between the thoracic and sternal ribs allow the size of the thoracic cage to be changed during inspiration and expiration. The resulting changes in pressure in the body cavity and air sacs allow air to be moved into and out of the lower respiratory system.81 Sternum of an adult chicken. Left lateral view: (a) keel (carina): (b) articular facet for coracoid bone: (c) rostrum; (d) cranio- lateral process; (e) articular facets for ribs; (f) thoracic pro- cess; (g) caudolateral process: (h) median trabecula; (i) lateral notch; (j) medial notch. The well: developed keel is characteristic of all ‘carinates’. In “ratites’ sueh as the ostrich on the other hand, the Keel is flat and raft-like. The keel provides attachments for the strong flight muscles and is especi- ally prominent in good fliers The lateral and medial notches are closed in life by fibrous mem: branes and with the thoracic and ccaudolateral processes strengthen the body wall. In the young bird the caudal part of the median trabecula is cartilaginous, flexible. In the pi lateral process arises separ from the thoracic process and fuses caudally with the median trabecula, The thoracic process is absent in the duck and goose. ‘The cranial part of the ster in the Whooper Swan (Cygnus eygnus) is. excav cavity in which the coils of the enormously long trachea lie fated to form a girdles of an adult fal view, In birds the thoracic girdle consists of the scapula, clavicle and. coracoid bones. The blade-like scapula (x) is relatively immobile and is firmly held to the ribs by muscle and ligaments, Cranially it articulates with the clavicle, coracoid bone and humerus. The extremely large coracoid (b) extends between the sternum and the shoulder joint. It acts to hold the wings away from the sternum during flight, and with the ribs prevents the thorax collapsing during the downstroke of the wing. The right and left clavicles (c) fuse ventrally to form the furcula (wishbone). They are absent or very reduced in some species of parrot. The furcula acts as.a spring holding the shoulder joints the optimum distance apart during wing movement83 Left sternocoracoclavicular membrane of an adult chicken, This tough, fibrous sheet (a) extends between the clavicle (b), coracoid (e) and. sternum (d) and anchors the three bones together. 84 Triosseal canal of an adult chicken. Where the clavicle (a), coracoid (b) and scapula (c) meet ‘medial t0 the shoulder joint is a space (arrow). the triosseal canal. Through the canal runs the tendon of the supracoracoid muscle which raises the humerus and causes the upstroke of the wing, ‘85 Left humerus of an adult chicken with the wing. rotated away from the trunk. The shoulder joint (2) is formed between the humerus (a), scapula (b) and coracoid bone (c). Movements of the humerus include elevation, depression, protraction and retraction. There is als ‘movernent around the axis of the wing including dorsal rotation which elevates the leading edge of the wing and ventral rotation which depresses the leading edge of the wing. The pectoral muscle. which is the main depressor of the humerus and therefore of the wing, attaches to the pectoral crest (2). When the wing is folded the humerus lies against the thoracic cage parallel to the86-88 Breast light) museles of an adult chicken. On each side there are two flight muscles, a superficial pectoral muscle (a) and a deep supracoracoid muscle (b). The pectoral muscle extends from the keel ofthe sternum, the clavicle and the sternocoracoclavicular membrane to the Pectoral crest of the humerus (sce 85) and is the main depressor of the wing. The supracoracoid muscle extends from the keel of the sternum through the triosseal canal (arrow) (see also 84) to the dorsal surface of the humerus ané is the main elevator of the wing. In adult turkeys and broiler breeders the supracoracoid muscle is the site of Oregon muscle disease (deep pectorel myopathy). The condition arises after sudden excessive activity and the accumulation of interstitial fluid in the muscle. Because the fascial coat ofthe musele prevents it from expanding, the accumulating interstiial uid presses onthe blood vessels causing foci of necrosis to develop. The breast muscles of the chicken and other species with a rapid jump take off contain more white muscle fibres than red fibres: because white fibres have relatively small " amounts of myoglobin, the breast muscles in these species are pale. White muscles are characterized by being Powerful but they are not suited for sustained activity. The breast muscles can be used for intramuscular injections. However. the needle should be inserted into the caudal part of the muscle rather than the cranial part where the blood supply is well-developed and there is greater chance of injecting into a vein. 186 Right pectoral muscle of an adult chicken, ‘88 Left supracoracoid muscle of an adult chicken, 87 Left supracoracoid muscle of an adult chicken ‘with the pectoral muscle reflected,89 Breast muscles of an udult pigeon, Ventral View. The breast muscles in the pigeon are red because they contain predominantly red fibres. Red muscle fibres use fat rather than glycogen as an energy source. Consequently they are more efficient than white fibres, because fat yields more energy than does carbohydrate per unit weight. Muscles which are used for sustained activity, therefore, like the breast muscles of good fliers. contain a high proportion of red fibres. 90 Gross anatomy of an avian long bone. As shown in this section of the humerus of a large seabird the cortex (a) is exceptionally thin and the medullary cavity (b) contains o network of trabeculae which increases the strength of the bone, The thin and brittle nature of the outer shell treatment of fra while intramedullary pinning would have the effect of destroying the internal strutting. Many of the bones of the post-cranial skeleton are pneumatized by diverticula of the air sacs which invade the medullary cavity of the long bones and the medullary spaces of the spongy bones where they replace the bone marrow. In female birds. prior to the laying season, a cancellous type of bone, medullary bone, is iaid down in the medullary cavity of the long bones under the influence of ‘oestrogens and androgens. During the egg laying eycle this bone is clternately undergoing formation and destruction and, along with the diet, acts as a source of calcium for the 91 Left forearm bones of an adult chicken with the ‘wing rotated away from the trunk. In the chicken the ulna (a) is stouter than the radius (b), the two bones being bowed along their length so that they are protected against bending forces in the plane of the wing. Attaching to the ulna by ligaments are the roots of the secondary flight feathers (see 39). When the wing is folded the ulna and radius lie close to the thorax and parallel to the humerus. The elbow joint (1) is formed by the condyles of the humerus (c) and the proximal ends of the radius and ulna. Movements at the joint are principally flexion and extension in the plane of the wing surface, with some rotation about the long axis of the forearm. At the carpal joint (2) the ulna and radius articulate with the ulnar (4) and radial (e) carpal bones. ‘manufacture of the egg shell. The presence of medullary bone increases the weight of the chicken skeleton before laying by approximately e\/ 41‘92 Left manus of an adult chicken with the wing rotated away from the trunk, By the second or third week after hatching the distal row of carpal bones has fused with the metacarpus to form the ‘carpometacarpus (a). The carpometacarpus articu- lates at the wrist with the radial and ulnar carpal bones (b). The three digits are the alular digit (c) with two phalanges, the major digit (d) with two phalanges, and the minor digit (e) with one phalanx. Attaching to the carpometacarpus and the phalanges of the major and minor digits are itium; (c) isehium; (¢) pubis: (e) caudal part of the synsacrum; (f free caudal vertebrae. (1) Tioischiadic foramen; (2) obturator foramen; (3) ischiopubic fenestra. The avi consists on each side of partly fused. ilium, ischium and pubis, the ilium being also joined to the synsacral part of the vertebral column (see 78). Except in a very few species the pelvic girdle is incomplete ventrally. The ilioischiadic foramen e. The large size and arched shape of the pelvic girdle and its fusion the primary flight feathers (see 39). The alular digit forms the skeleton of the bastard wing (see 43) which during slow flight is extended aw from the wing to form a slot preventing stalling. Amputation or surgical pinioning of the wing to Prevent escape in waterfowl and other ‘ornamental birds is performed by cutting off the manus of one wing through the proximal part of the carpometacarpus. so. that the bird is off-balance when tying to fly. The optimum age for the operation is in the down chick stax Surgical pinioning is illegal in birds kept for agricultural purposes. Movements at the wrist joint are mainly flexion and extension in the plane ‘of the wing surface and are coupled to movements ‘of the elbow joint. Thus, during flexion of the elbow joint the wrist i flexed while extension of the elbow joint causes the wrist to be extended. Movements of the digits include flexion. ‘extension, elevation and depression. Extension of the major and minor digits has the effect of spreading the primary flight feathers, while hes the effect of folding the primary with the vertebral column are adaptations for supporting the trunk in the bipedal standing position. Aviculturists sometimes try to sex birds by assessing the distance between the pubic bones, the supposition being that the bones are further apart in females than in males. However, it is generally accepted that the test is subject and unreliable, The space between the pelvi bones is also used to establish whether a pullet in lay o not. In the so-called ‘finger’ test, two fingers comfortably inserted between the Dones is. ‘an indication of a laying hen, while three fingers is ‘an indication of a bird heavily into lay.94 Right femur of an adult chicken. Lateral view The distal end of the femur (a) slopes craniolaterally bringing much of the hind limb close to the centre of gravity of the body. Within the single joint cavity of the hip joint (1) are two articulations, one formed between the head of the femur and the acetabulum of the pelvic bone as in ‘mammals, and one formed between the femoral ‘trochanter (arrow) and the antitrochanter of the ilium (see 78). Movements of the thigh include protraction, retraction, abduction, adduction and rotation. As well as rotating the limb, the rotatory muscles are responsible during locomotion for bringing the body over the supporting leg when, the opposite leg is in the air, which results in the ‘waddling type of gait typical of birds with short legs such as ducks. The knee joint (2) is formed between the distal end of the femur, the patella (b), and the proximal ends of the tibiotarsus (c) and fibula (d). Its main movements are flexion and extension. 95 Ischiadic nerve of a chicken. In. this photograph the position of the femur is indicated by the muscle mass (a) which covers its medial face. The ischiadic nerve (b) lies behind the id below two muscles on the medial aspect of the limb which have been removed Marek’s disease is a condition of poultry in which there is a proliferation of lymphoid tissue and a special predilection for peripheral nerves which become irregularly thickened. The ischiadic nerve is one of the nerves which is usually examined post-mortem. Other nerves which are inspected are show in 193, 194, 196, 230, 231, 233 and 269. While intramuscular injections ean be made into the thigh muscies, there is a danger that the ischiadie nerve may be damaged, Furthermore, substances injected into the hindlim’ may pass through the renal portal system (see 195) before entering the systemic circulation,96, Right craral bones of an adult chicken. Lateral view. This part of the limb is frequently referred to.as the ‘drumstick’ and consists of the splint-like fibula (2) and the tibia which has fused with the proximal row of tarsal bones to form the ubjotarsus (b). However, the proximal row of tarsal bones remains free for the first few months after hatching. The cranial cnemial crest (arrow) provides an attachment for the main extensor muscle of the knee joint. Unlike the other long bones, the tibiotarsus and femur are good sources of bone marrow. 97 Right tarsal (hock) region of an adult chicken. Dorsal view, The articulation is an intertarsal joint between the tibiotarsus (a) and the tarsometatarsus (b). The main movements are flexion and extension. On the cranial surface of the distal part of the tibiotarsus is the extensor canal (arrow) through which the tendons of the extensor muscles of the toes pass.‘of an adult chicken opened 10 show the tarsometatarsal articular surfaces. In the lateral part of the joint is cartilaginous menis (arrow) which lessens the incongruence of the articular surfaces. A medial meniscus is absent in the chicken. The hock joint is routinely examined in the poultry slaughter plant cartilage of an adult chicken. Fixed preparation: (a) tibiotarsus: (b) hock joint; (c) tibial cartilage; (d) tarsometatarsus, The tibial cartilage is a fibrocartilaginous block on the caudal aspect of the hock joint, Passing over it are the tendons of the gastrocnemius and superficial digital flexor muscles (1) while the tendons of the deep digital flexors (2) travel through canals in the cartilage. In the chicken and turkey ossification may develop in many of the tendons of the wing and hindlimb. Bone formation in the chicken starts after about 90 days and occurs especially in the tarsometatarsal region and on. the caudal tibiotarsus, dorsal carpometacarpus and caudal antebrachium, 45101 Feet of an adult chicken: (a) tarsometatarsal bone; (b-c) digits I-IV. The tarsometatarsus is formed by the fusion of the distal row of tarsal bones and the metatarsal bones of digits II, II] and LV. Evidence for this is visible at its distal extremity which hes three articular trochleae. A bony process (arrow) arises fom the distal part of the medial surface in males and many females and is the core of the spur (sce 19), A small movable metatarsal I remains separate and is attached to the tarsometatarsus by ligaments. Digits F-1V have two, three, four and five phalanges 46 100 Caudal tibial vein of a 10-week-old chicken. The caudal tibial vein (arrow) crosses the medial aspect of the tarsal joint where it is casily accessible. Except in the smallest species it can be used for intravenous injections. respectively. This seems to be a general Phalangeal formula for birds. The most distal Phalanx forms the bony core of the keratinized claw. When a bird crouches during perchin flexion of the hock joint tenses the tendons which cross the caudal surface of the joint and passively clamps the digits around the perch. On rising the hock is extended which Temoves the tension from the flexor tendons, so that the digits can be extended. This mechanism should be remembered when attempting to undo the grasp of a large birdDigestive system 102 Oral cavity and pharynx of an adult chicken. Because a soft palate and oropharyngeal isthmus are absent, the oral and pharyngeal cavities together form a common cavity which is often referred to as the ‘oropharymx’. On the palate (a) is the choana ((), the median fissure opening into the nasal cavi- ties. The infundibular cleft 2) is the common open- of the auditory tubes. The tongue (b) in the chicken, turkey and pigeon cannot be protruded ‘and is mainly an organ of swallowing, The laryngeal mound (c) has on its rostral surface the glottis G3) is the entrance to the larynx. Many backward- pointing, thickly keratinized papillae (arrows) are distributed on the roof of the oropharynx, on the tongue and on the laryngeal mound. The opening Of the mouth is generally called the gape. Swallow- ing is achieved by rapid rostrocaudal movements of the tongue and the laryngeal mound assisted by much sticky saliva and the caudally-directed papillae which propel the food backwards and prevent its regurgitation. During this process the choana, in- fundibular cleft and glottis are reflexly closed. The ‘method of drinking varies with the species. In the chicken water is propelled into the mouth by rostro- caudal movements of the tongue. The head is then raised and the water passes into the oesophagus mainly by gravity. In the pigeon, in contrast, the bbeak is immersed in the water throughout and the head is not raised. 103 Palate of an adult chicken. Ventral view: (a) choana; (b) papillae. The numerous openings of the mucus-secreting salivary glands which form an almost continuous jayer in the walls of the oropha- rymx are indicated by arrows. (Scale, J mm) a105-108 Tongue of a duck. The tongues of ducks. geese and swans are adapted along with the lamellae Of the bill either for straining minute organisms from water or for cropping arass and weeds. The fleshy tongue of the domestic duck is a straining ‘organ. Its rostral part has a scoop-like process (a) ‘and the lateral margin has a double row of overlapp- 108 Tongue of an adult duck. Dorsal view 104 Hyoid apparatus of an adult chicken. Dorsal view: (a) entoglossal bone: (b) rostral basbranchial bone; (c) caudal basibranchial bone; (d) cerato- branchial bone: (e) epibranchial bone. The ento- slossal bone lies in the body of the tongue and the rostral basibranchial bone in the base of the tongue The hyoid apparatus and its associated musculature are responsible for tongue movements. Except in parrots, there are no true intrinsic muscles in the tongue. (Seale, 5 mm) ing bristles (b) which are interspersed caudally with tooti-like processes (c). The bristles interdigitate witha double row of lamellae (d) on the bill. On the dorsal surface of the tongue is a shallow median groove (¢) rostrally and a fleshy cushion or eminence (atthe root. 106 ‘Tongue of an adult duck. Dorsal view. (Seale, 2mm)—_— os 107 Tongue of an adult duck. Lateral view 108 Tongue of an adult duck. Lateral view. (Scale 109 0 Sei doo aerate wine toe net Bi te Nae { the combined auditory tubes. The rost chicken there are numerous lize (arrows) on the tor udally-directed p: and lacy 'b) which help propel the food backwards into the esophagus. of the palate has fleshy, vascular cushions >) which are used with the tongue to hold food. aLIL Neck of an 8-week-old chicken, Ventral view The thin-walled oesophagus (a) lies for most of its course in the neck immediately below the skin and to the right of the trachea (b). At the thoracic inlet it enlarges ventrally to form the crop (c). (d) Lobes of thymus. 112 The relationship between the crop and skin in aan adult chicken. An area of skin on the right side of the thoracic inlet has been removed to show its close attachment (arrows) to the crop (a). Part of this attachment is to two sheets of striated muscle which appear to form sling-like supports for the diverticulum, When full the crop can easily be palpated in the live bird, 113 Interior ofthe oesophagus and crop of an adult chicken. The crop (a) is a ventral diverticulum of the oesophagus (b), which it divides into cervical and thoracic parts. Because the internal surface of the oesophagus is longitudinally folded (arrow). it is relatively distensible114 Eull crop of an adult chicken. Ventral view ‘The function of the crop (a) is 10 store food whe! the gizzard compartment of the stomach is full. th food in the crop undergoing softening and swelling in preparation for chemical digestion in the stomach and intestines. When the gizzard is empty food by- passes the crop and travels directly 10 the stomach, the entrance to the crop being closed off. In the evisceration of poultry in the slaughter plant the oesophagus (b) is cui below the crop so that to avoid contamination of the carcass by food the bird is not fed for 2 to § hours before death 115 Light micrograph of a section through the entire ‘wall of the oesophagus of a chicken. Masson's crich: rome: (a) partly Keratinized stratified squamous epithelium: (b) lamina propria; (c) mucous glands (a) muscularis mucosae: (e) submucosa; (f) inner circular muscle layer; (g) outer longitudinal muscle layer. The structure of the crop resembles that of the ocsophagus, except that the mucous glands are restricied to an area close to the oesophagus. (Seale, 250 um)116 Ocsophagus and crop of an adult budgerigar. Ventral view. The cervical oesophagus (a) lies mai: ly to the right of the trachea (b); at the thoracic inlet it expands to form the crop (c), which, as in other psittacine species, is stretched transversely across the neck. Seed from the erop (or proventri- clus) is regurgitated by budgerigars to feed their young, Food may also be regurgitated by the male as pari of courtship behaviour. Regurgitation can even occur when a budgerigar is alone, in response either toits own image reflected in a mirror, or toa toy. oF to the appearance of the owner 117 Highly developed crop of an adult pigeon. Ven- tral view. As in the chicken the crop is a ventral diverticulum of the oesophagus (a), but unlike in the chicken itis divided into two large lateral sacs (b). The crop in both sexes of adult pigeon produc ‘crop milk” which is regurgitated and fed to the young. during the first few days after hatching. The ‘milk’ is a desquamation of fat-laden cells of the prolife ated stratified squamous epithelium lining the crop and its manufacture is controlled by the pituitary hormone prolactin. While it resembles mammalian milk, it has no carbohydrates or calcium, 118 Crop of an adult canary. Ventral view. The ‘oesophagus (a) lies to the right of the trachea (b). and close to the thoracic inlet. It forms a spindle shaped swelling which represents the crop (€). 119 Qesophagus and crop of an adult duck. Ventral view, The oesophagus (a) close to the thoracic inlet. forms a relatively indistinct spindle shaped widen ing (b) which represents the crop. In the duck the wall ofthe terminal part of the thoracic oesophagus reyated Iymphoid nodule which is commonly referted to as the ‘oesophageal tonsil120 Ventral view of the thorax and abdomen of un adult chicken with the sternum and abdominal wal removed. Few viscera ate visible because the abdo- minal contents are covered ventrally by a double- layered fat laden fold of peritoneum, the posthepa tie septum (a). The only organs which are not cove- red are the right and left lobes of the liver (b) and the heart (6). Part of the gizzard (arrow) can be seen suspended in the postnepatic septum. Note that in birds no diaphragm separates the thorax from the abdomen, 122 Stomach of an 8-week-old chicken. Right la al view. The stomach consists mainly of a cranial glandular compartment or proventriculus (a). and caudal muscular compartment or gizzard (6) Between the two compartments is the intermediate zone which in the domestic birds is marked on the outside by a constriction, the isthmus (c). The pylo- ric part of the stomach (d) arises from the right face of the gizzard and connects the gizzard with the duodenum. The stomach of the chicken, like that of the other domestic birds, is characteristic of species which feed on relatively indigestible types of food that have to be physically broken down prior to chemical digestion. Typically in these birds the gizzard is highly developed with a very thick muscular coat, and the proventriculus and gizzard are clearly set off from one another, ray 121 Ventral view of the thoracoabdominal cavity of the adult chicken sown in 120, but with the posthe- patie septum removed. On the left side ofthe abdo- men lies the gizzard (a) and on the right side lies the narrow U-shaped loop of the duodenum (b). In all species ths is the most ventrally-situated part of the intestinal tract. Although the gizzard is usually covered by a fat-laden posthepatic septum, it can still be palpated. except in the smallest species, in the live bird. The heart (d) is clearly visible above the liver (C) 12123 Interior of the proventriculus of an adult chick- fen, Projecting into the lumen are welldeveloped papillae (arrows) at the tops of which open the compound gastric glands. The glands form a thick layer (a) in the wal of the proventriculus, Gastric prozcalysis occurs principally in the gizeard where the pHs low (1.5-2.5). 124 Light micrograph of a seetion through the entire wall ofthe proventriculus of a chicken. Masson chrome: (a) columnar mucus-seereting epithelium; (b) lamina propria; (c) multilobular glands of the lamina propria: (d) muscularis mucosae; (e) submu- cosa: (f) inner circular muscle layer; (g) outer long tadinal muscle layer. ‘The muscularis: mucosa surround the glands of the lamina propria, The glands consist. mainly of closely-packed alveoli radiating out from a central cavity. The central Cavity is drained by a secondary duct, all the secon- Gary duets ofa gland uniting to form a primary duct ‘which opens into the lumen of the organ. The glan- ‘ular epithelium of the alveoli is lined mainly by one type of cell, the oxynticopeptic cell, which secretes both pepsin and hydrochloric acid. (Scale, 500 wn)125 and 126 Gizzard of a chicken. Most of the wall of the gizzard is made up of smooth muscle rich in ‘myoglobin and arranged into four semi-autonomous masses. These are the dark-coloured caudodorsal (1) and cranioventral (2) thick muscles. and the lighter-coloured craniodorsal (3) and caudoventral 125 127 Interior of the stomach of an adult chicken. Separating the proventriculus (a) and gizzard () is, the intermeciate zone (b) which is characterized by 4 histological structure intermediate to that of the proventriculus and gizzard. Compound glands are absent here and the internal surface is relatively smooth. The bulk of the gizzard is formed by the body (1). in the walls of which are the thick muscles (2). At either end of the gizzard are the cranial (3) and caudal (4) blind sacs, in the walls of which are the eraniodorsal and caudoventral thin muscles (3). The entrance (6) to the pyloric part of the and the duodenum lies in the cranial r surface of the gizzard is a hardened membrane, the cuticle (sometimes called the koilin layer). which is a carbohydrate-protein complex secreted by the gizzard glands. The men brane is especially well-developed in birds, such as the domestic species, which feed on relstively hard items of food. In these birds it is asymmetrically developed, being thickest opposite the thick muse les of the body of the gizzard and thinnest opposite the thin muscles of the cranial and caudal blind sacs. Its green or yellow colour is due to bile reg gitated from the duodenum via the pylorus. (4) thin muscles. The muscles 2 ich to the right and leit tendinous centres (5) in the lateral walls of the gizzard. Because the muscles are asymmetrically arranged, rotatory as well as crushing movements are set up when the gizzard contracts, }-week-old chicken. Right lateral 127129 and 130 Histological structure of the chicken sgizaard: (a) cuticle; (b) glands in the mucous mem- mucosa; (d) muscle. The cuticle is & carbohydrate-protein complex secreted by the tis composed of vertical rods (arrows) and horizontal brane: (c) st mucosal glands and the surface epithelium. matrix. The vertical rods are secreted by the muco- 29 129 Light micrograph of a section through the entire wall of the gizzard of a chicken. Masson’s trichrome. (Scale, 250 um) 36 128 Gizzard cuticle of an adult chicken. The cuticle is a secretion of the mucosa and can therefore be easily stripped away from the organ. In the poultry slaughter plant the cuticle has to be removed by machine before the gizzard is added to the giblets. sal glands, the sceretion hardening within the lumen. as a thickening perpendicular to the surface, The horizontal matrix between the rods is a secretion of the surface epithelium which hardens after spreud- ing around the rods. Within the horizontal matrix are trapped desquamated cells of the surface epith- lium, 130 Light micrograph of the inner layers of the gizzard of a chicken. Masson's trichrome. (Seale, 100 um)132 131 Gizzard contents in an adult chicken. Large amounts of grit (arrows) are mixed up with the food. Grit is also ingested by the other domestic species, because like the chicken they feed on rela tively coarse material which must be reduced before chemical digestion, The need to take in grit has led to problems in the Mute Swan (Cygnus ofr) which frequently mistakes the small lead weights used by anglers to sink their fishing lines as a form of grit and ingests them. The weights are slowly eroded in the gizzard and the resulting lead salts absorbed in the small intestine to cause lead poisoning. In this, photograph the three factors which are important in the physical phase of gastric digestion in the domestic birds are shown, They are the grit in the lumen, the hardened cuticle lining (a), and the massively developed semiautonomous muscles (b) 132-134 Spleens. In all birds the dark-red spleen (a) lies on the right side of the junction between the proventriculus (b) and gizzard (c). The shape varies from spherical in the chicken, to triangular in the 132 Spleen of a 6-week-old chicken. 134 duck, to elongated in the pigeon and canary. While having a similar function to that of mammals, the spleen does not appear to be important in storing blood. 133 Spleen of an adult duck. 134 Spleen of an adult pigeon.135 Duodenum and pancreas of an adult chicken. ‘Ventral view. The duodenum is arranged as a narrow U-shaped loop with descending (a) and ascending (b) parts, the two parts being held closely together by mesentery. The duodenal loop encloses the pan- creas (c). It is always the most ventral part of the intestinal tract (see 121). Within the pale red or yellow pancreas of the chicken dorsal, ventral, third and splenic lobes have been identified. 136 Duodenal loop of an adult canary. Ventral view. As in otlier avian species the duodenal loop (a) is the most ventrally situated part of the intestinal tract and encloses the pancreas (bh). Compared with the narrow duodenal loop of the chicken, however. that of the canary is extremely broad. See also the sizzard () 137 Liver of a S-week-old chicken, Ventral view. In birds the liver (a) lies in the cranial part of the thoracoabdominal cavity, its cranioventral portions surrounding the apex of the heart. It consists of right (1) and left (2) lobes, the two lobes in the chicken being almost equal in size. The left lobe in the chicken and turkey is subdivided into lateral (3) and medial (4) parts. In large birds it may be possi- ble to palpate the liver beyond the edge of the ster- num, The edges of the liver are normally thin and sharp but if the liver is enlarged they may become rounded. The colour of the liver depends on the nutritional state of the bird and, while itis frequent- ly red-brown as in the photograph, it may be light brown or, ifthe bird is on a high-fat diet, yellow. “The caudal vena cava (b) passes through the cranial part of the right lobe and receives the hepatic veins. ‘The gall bladder (c) lies on the viscerel surface of the right lobe. In the poultry slaughter plant the gall bladder is frequently cut by accident during post-mortem processing and this is a common cause of careass contamination. The liver, but not the gall bladder, forms part ofthe giblets. 38138 Liver of an adult pigeon. Ventral view. In this species, ay in the duck and goose. the right liver lobe (a) is very much larger than the lft (b), and the left lobe is not subdivided, A gall bladder is absent in the pigeon snd budgerigar. It is pres however, in some species of parrot. Heart (C) 140 Mucous membrane lining the distal part of the jodenum of an adult chicken. The papilla on which the pancreatic and bile ducts open is indicated by an arrow. (Scale, 2 mm) 139 Pancreatic and bile duets of an adult chicken. The pancreatic (+) and bile (2,3) duets open into the distal part of the ascending limb of the duode- um (b). In the chicken the common hepatoenteric uct (2) drains the right and left lobes of the liver and is formed by the union of the right and left hepatic ducts on the visceral surface of the right lobe. The right hepatic duct gives off a branch which enters the gall bladder. The gall bladder is rained by the eysticoenteric duet (3). In the pigeon, which has no gall bladder. there are two hepatoen- tetic ducts, the right duct opening into the middle of the ascending limb of the duodenum and the better developed left duct opening into the proxi- ‘mal part of the descending limb, See also descending limb of the duodenal loop (a), pancreas (¢), liver @ if * ai 140143 Yolk sac of a day-old chicken to show the normal appearance of yolk, 144 Liver of a day-old chicken. Ventral view. At hatching the liver lobes (arrows) have a bright yellow colour because of the pigments they absorb ‘along with the lipids of the yoik. The colour gradu- ally changes to the mahogany-brown of the adult liver between day’ 8 and I+. oo 141 Jejunum and ileum of a 6-week-old chicken. The jejunum and ileum (a) in the chicken and turkey are relatively uncompli cated, unlike in most species of birds, and form short, garland-like coils at the edge of the long dorsal mesentery. Within the me- semery are the cranial mesenteric artery and vein and their branches (b), 142 Yolk sac of a 3-day-old chicken. The yolk sac (a) and yolk duct (b) open into the antimesenteric side of the small intestine (©) opposite the terminal branches of the cranial mesenteric artery. They are used arbitrarily to distinguish the jejunum prox- imally from the ileum distally. About 10 days after hatching, the residual yolk has been absorbed into the blocdstresm either through the wall of the sac or through the wall of the gizzard via the small intestine When empty, the volk sac isconverted into sear tissue.